Report on a collection of amphibians and reptiles from Sichuan, China

UNIVERSITY OF

ILLINOIS LIBRARY

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BIOLOGY

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NEW SERIES, NO. 58 a0O>

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Report on a Collection of Amphibians
and Reptiles from Sichuan, China

Robert F. Inger
Ermi Zhao
H. Bradley Shaffer
uanfu Wu

July 31, 1990
Publication 1413

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

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Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford University Press, Stanford, Calif., 943 pp.

Grubb, P. J., J. R. Lloyd, and T D. Pennington. 1963. A comparison of montane and lowland rain forest in Ecuador.

I. The forest structure, physiognomy, and floristics. Journal of Ecology, 51: 567-601.
Langdon, E. J. M. 1979. Yage among the Siona: Cultural patterns in visions, pp. 63-80. In Browman, D. L.,and R. A.

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FIELDIANA

Zoology

NEW SERIES, NO. 58

Report on a Collection of Amphibians
and Reptiles from Sichuan, China

Robert F. Inger Ermi Zhao

Department of Zoology Chengdu Institute of Biology

Field Museum of Natural History Chengdu, Sichuan
Chicago, Illinois 60605-2496 Peoples Republic of China

H. Bradley Shaffer Guanfu Wu

Department of Zoology Chengdu Institute of Biolog

University of California at Davis Chengdu, Sichuan

Davis, California 95616 People's Republic of China

Accepted September 16, 1988
Published July 31, 1990
Publication 1413

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 1990 Field Museum of Natural History

ISSN 0015-0754

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

Abstract 1

Introduction 1

Materials and Methods 2

Species Accounts 5

Summary of Habitat Distribution of

Anurans at Wa Shan 22

Acknowledgment 23

Literature Cited 23

List of Illustrations

1 . Sichuan Province, showing sample sites 2

2. Vegetation at Wa Shan, Sichuan 3

3. Vegetation at Qiliba, Sichuan 4

4. Oreolalax rugosus (Liu) 8

5. Calluella yunnanensis Boulenger 11

6. Rana daunchina Chang 13

7. Southern Sichuan and adjacent prov-
inces showing localities of samples of
Rana yunnanensis 14

8. Polypedates chenfui (Liu) 15

9. Achalinus meiguensis Hu and Zhao .... 17
10. Amphiesma metusia sp. nov 19

List of Tables

1. Comparison of Qiliba and Wa Shan sam-
ples of adult male Bufo andrewsi 9

2. Comparison of body proportions in
males of Rana chaochiaoensis and R. ja-
ponica from Qiliba and Wa Shan, Si-
chuan 10

3. Perching sites of frogs at Wa Shan, Si-
chuan 21

4. Distribution of stream-breeding frogs
along streams of various widths at Wa
Shan, Sichuan 22

5. Distribution of size classes of Rana bou-
lengeri along streams of various widths at
Wa Shan, Sichuan 23

Report on a Collection of Amphibians
and Reptiles from Sichuan, China

Abstract

A collection of amphibians and reptiles was made
in Sichuan Province, P.R.C., in 1987 by a team
from Chengdu Institute of Biology and Field Mu-
seum of Natural History. At the principal site,
forested slopes and top of Wa Shan 40 km WNW
of Mt. Emei, 18 species of amphibians and 12
species of reptiles were collected between 1 000 and
2520 m in four weeks. Twelve species of amphib-
ians and one reptile were collected at Qiliba, about
240 km S of Wa Shan, between 2950 and 3 140 m
in five days. One new species of snake, Amphiesma
metusia, is described. The amphibian fauna at Wa
Shan shows ecological segregation based on types
of breeding sites and perches and on altitudinal
distribution.

Introduction

In 1987 Chengdu Institute of Biology and Field
Museum of Natural History initiated a joint pro-
gram of field studies on the herpetofauna of Si-
chuan, thus renewing an old association that had
led in 1950 to the publication by Field Museum
of "Amphibia of Western China" by Liu Qeng
Qao. The current program began with a period of
collection and observation by the present authors
from May 21 to July 3, and the first results are
summarized in this paper.

We chose to begin this program with an inten-
sive study of microhabitat distribution of am-
phibians and reptiles at one site, as previous field-
work in Sichuan has concentrated on elucidating
the geographic distribution of the fauna. The site
' ultimately chosen was Wa Shan (29°38'N,

102°56'E), a mountain 40 km WNW of Mt. Emei
which had been explored herpetologically by Liu
and his associates. The field party worked at Wa
Shan for four weeks, 22 May-1 6 June. While look-
ing for an appropriate site, we visited Qing Cheng
Shan (76 km NW of Chengdu in Guan Xian) and
collected there for two days. After leaving Wa Shan,
we worked five days at Qiliba (27°53'N, 102°34'E),
35 km E of Xichang City, which is 500 km S of
Chengdu. Localities are shown in Figure 1 .

Most of the collecting at Wa Shan was carried
out between 1 200 and 1 500 m. The steepest slopes
throughout and the less steep ones around 1400
m were covered with evergreen angiosperm forest
having dense undergrowth of herbs, ferns, and
shrubby angiosperms. This original forest is being
actively logged. As areas are cleared, conifers are
planted. Below 1 300 m the flatter areas have stands
of these conifers, some as large as 30 cm DBH.
There is little undergrowth beneath the conifers,
although needles and dead branches cover the soil.
The groves of conifers meet angiosperm forest at
places, particularly along small, steep watercours-
es, to form patches or strips of mixed forest. On
some slopes seedling conifers had just been plant-
ed, leaving these areas as clearings. The largest
stream (ca. 8 m wide) in our working area was
flanked in spots by flat patches supporting grass,
bamboo, and shrubs. We also collected along sev-
eral small streams (1-2 m wide) flowing through
angiosperm forest. We spent two days collecting
in a conifer-rhododendron forest on the rather flat
top of Wa Shan at 2520 m (fig. 2).

Qiliba is on an undulating plateau 2500-3200
m ASL with marshy grassland in flat areas and
chaparral-like, shrubby vegetation (fig. 3) on steep-
er slopes. Scattered small trees occur on some
slopes. Small streams (1-3 m wide) are common,

<H

GER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

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Fig. 1. Sichuan Province, People's Republic of China. 1, Qing Cheng Shan; 2, Wa Shan; 3, Qiliba.

and ponds (2-10 m in diameter) dot the fiat areas
(fig. 3).

Materials and Methods

The location of each specimen was recorded fol-
lowing the habitat classification scheme described
elsewhere (Inger & Colwell, 1978), modified to
include the vegetation types encountered in Si-
chuan. Each specimen was bagged separately as
captured and usually preserved in formalin within
three hours. With few exceptions, all specimens
from Wa Shan were weighed to the nearest gram
prior to preservation. Snout-vent length (SVL) and
total length were measured for snakes before pres-
ervation. Lizards and amphibians were measured

after preservation. When the number weighed is
less than the number measured, the former is given
in parentheses. Denticle counts of tadpoles follow
the system used by Liu (1950) and Liu and Hu
(1961).

For testing statistical significance of ecological
distributions in the last part of this paper (Sum-
mary of Habitat Distribution of Anurans at Wa
Shan), we used chi-square tests, mostly in the for-
mat of 2 x 2 tables in which occurrence of a given
species in a particular type of microhabitat was
compared against that of "all others." Synonymies
are presented in the following sequence: original
description, first use of present combination (if
different from original), synonyms.

The collection has been divided, half deposited
in Chengdu Institute of Biology and half in Field
Museum of Nature History.

FIELDIANA: ZOOLOGY

Hg. 2. Vegetation at Wa Shan, Sichuan. Above, broad-leaved evergreen forest at about 1400 m; below, pine,
rhododendron, axalea forest at 2520 m.

INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

Fig. 3. Vegetation at Qiliba, Sichuan. Above, chaparral at 3020 m; below, wet meadow flanked by chaparral at
3140 m.

FIELDIANA: ZOOLOGY

Species Accounts

CAUDATA

HYNOBIIDAE

Batrachuperus pinchoni (David)

Dermodactylus pinchoni David, 1871, Nouv. Arch.

Mus. Nat. Hist. Nat., 7: 95. Type locality: Muping

(= Baoxing), Sichuan.
Batrachuperus pinchoni Stejneger, 1925, Proc. U. S.

Nat. Mus., 66: 5.
Batrachuperus longdongensis Liu and Tian, 1983, Acta

Zootaxon. Sinica, 8: 210. Type locality: Longdong,

Mt. Emei, Sichuan.

Four adult males, SVL 90-120 mm; tail length/
total .43-.49 (mean .459); weight 21-25 g (N = 3,
mean 23.7); head width/SVL . 1 5-. 1 8 (mean . 1 72).
Two subadult males, SVL 60-61 mm; tail length/
total .46-.47; weight 7 g (N = 2). Five adult fe-
males, SVL 90-1 14 mm; tail length/total .45-.48
(mean .465); weight 18-27 g (N = 2); head width/
SVL . 1 6-. 1 9 (mean . 1 73). Four subadult females,
SVL 55-72 mm; tail length/total .44-.47; weight
6-10 g. Sixteen juveniles, no external gills or gill
slits, SVL 31-53 mm. Five larvae with external
gills, SVL 20.3-31.5 mm. Thirty-two from Wa
Shan, 3 from Longdong, 1 from Qiliba.

The Longdong specimens, topotypes of Batra-
chuperus longdongensis, were collected at 1250 m
in the same spring- fed, rocky, 10 m stream from
which the types came; it is bounded by fields on
one side and young secondary forest on the other.
The Qiliba salamander was in a small (.5 m) grav-
el-bottom stream flowing through chaparral and
marshy grassland at 3140 m. All the Wa Shan
animals were in streams .5-2.0 m wide. Thirty
were collected at 2520 m, 29 in a fir forest and
one in marshy grassland. Two were in broad-leaved
evergreen forest at 1530 m. We found two egg
cases and the five larvae on the top of Wa Shan
(2520 m)May 31-June 1.

The original description of B. longdongensis, re-
ported only from the type locality within the range
of pinchoni, differentiated it on the basis of large
size (114-265 mm) and an unstated percentage
("most") of neotenic adults. Our three topotypes
are adults (total lengths 184-201 mm), without
gill slits or external gills. They fall within the size
range of the Wa Shan sample (total lengths 1 72-
213 mm). A few adults with gill slits have been
found at Baoxing, the type locality of pinchoni, in
recent years. We believe longdongensis is conspe-
cific with pinchoni.

ANURA
DISCOGLOSSIDAE

Bombina maxima (Boulenger)

Bombinator maximus Boulenger, 1905, Ann. Mag.

Nat. Hist., (7), 15: 188, pi. 13. Type locality: Tong

Chuan Fu (= Malong), Yunnan.
Bombina maxima Schmidt, 1927, Bull. Amer. Mus.

Nat. Hist., 54: 556.

Three males, SVL 42-54 mm (mean 50.0). Col-
lected at Qiliba.

These were caught in chaparral at 2950-3000
m. One frog was in a small permanent pond and
the others in grass.

PELOBATIDAE

Leptolalax pelodytoides oshanensis (Liu)

Megophrys oshanensis Liu, 1950, Fieldiana: Zool.,
Mem., 2: 197. Type locality: Mt. Omei (= Emei),
Szechuan (= Sichuan).

Leptolalax pelodytoides oshanensis Dubois, 1983, Al-
ytes, 2: 149.

Thirty males, SVL 26-30 mm (mean 27.8); one
female, SVL 31 mm. All were collected at Wa
Shan.

All individuals were caught at night in forest
(broad-leaved or mixed broad-leaved and conif-
erous) at 1200-1400 m on banks of small streams,
1-1.5 m (16 individuals), 2 m (14), and 3 m (1)
wide. Males were calling from gravel (9 individ-
uals), low herbs (8), rocks (7), and small dead sticks.

Dubois (1983) provisionally considered osha-
nensis to be conspecific with pelodytoides. Previ-
ously, he had made Leptobrachium minimum
Taylor a synonym of oshanensis (Dubois, 1981).
Because specimens from Vietnam, Laos, Thai-
land, and China had less webbing than the types
of pelodytoides, Dubois recognized these popula-
tions as constituting the subspecies, pelodytoides
oshanensis (Liu). Liu (1950) had stated that osha-
nensis completely lacked webbing. Only one of the
Wa Shan frogs has webbing at the base of the third
to fifth toes. One-third of adults we have examined
from the type locality of minimum (Doi Suthep)
and adjacent areas of Chiang Mai Province, north-
ern Thailand (fmnh 2 1 3933-38, 2 1 6077-83) have
webbing at the bases of the outer toes. Liu and Hu
( 1 96 1 ) show webbing at the bases of the outer toes
in Yunnan frogs they refer to pelodytoides (their

INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

fig. 5). Thus if there is a geographic break in this
character, it seems to be between Sichuan frogs on
the one hand and all those to the south on the
other.

Liu (1950) said that larval oshanensis had only
four rows of denticles (1:3 + 3) on the upper lip as
opposed to five or six in pelodytoides from Thai-
land (presumably based on Smith, 1917). Nine
larvae from Wa Shan (Gosner stages 25-38) have
1:2 + 2 (2 tadpoles), 1:2 + 3 (2), or 1:3 + 3 (5). Liu
and Hu (1961) reported upper denticular formulas
of 1:2+2 (2), 1:3 + 3 (8), and 1:4+4 (3) for larval
pelodytoides from Yunnan. Larvae from Chiang
Mai Province, Thailand (fmnh 212386, usnm
103402) have 1:3 + 3 (2, stages 26, 32) and 1:4+4
(stage 37). Again, the break in character-state dis-
tribution seems to lie between Sichuan (maximum
number of upper labial rows = 4) and the popu-
lations from Yunnan southward (maximum num-
ber of rows >4).

Liu and Hu (1961) illustrate differences in ser-
ration of larval beaks, oshanensis having finer ser-
rae than pelodytoides from Yunnan (cf. their figs.
4-5). We confirm this difference. The larvae from
Wa Shan have more finely serrated beaks than
those from Thailand. Again, the change in char-
acter-states takes place immediately south of Si-
chuan.

Liu (1950) said that males of oshanensis had
paired vocal sacs and thus differed from pelody-
toides. We have dissected six males from Wa Shan,
all having single vocal sacs. As Wa Shan is very
close (ca. 50 km) to the type locality (Mt. Emei),
we think this character of oshanensis needs rein-
vestigation.

The similarity of the Wa Shan series with adults
we have seen from Thailand in size, color pattern,
size and distribution of glands, and other external
features is striking and leads us to accept Dubois's
assessment of conspecificity of pelodytoides, osha-
nensis, and minimum. However, as we have shown
above, changes in character-states are such that
the subspecies pelodytoides oshanensis should be
restricted to Sichuan.

76 mm (mean 67.8); weight 17-36 g (mean 28.0).
Two juveniles, SVL 27, 48 mm. All from Wa Shan.
Eight frogs were collected 1200-1360 m and 12
at 1 400 m, 1 7 in broad-leaved forest and three in
mixed broad-leaved and coniferous forest. Three
were found at some distance from water, 1 5 in or
on the banks of small (1-2 m) streams and two on
the banks of a larger (8 m) stream.

Megophrys shapingensis Liu

Megophrys shapingensis Liu, 1950, Fieldiana: Zool.,
Mem., 2: 194. Type locality: Shaping, Opienhsien
(= Ebian County), Szechwan (= Sichuan).

One juvenile, SVL 52 mm. From Qiliba, on
bank of small (2 m) stream in chaparral.

Oreolalax omeimontis (Liu and Hu)

Scutiger omeimontis Liu and Hu, 1960, Scientia Sin-
ica, 9: 767. Type locality: Changshenggou, Mt. Omei
(= Emei), Szechwan (= Sichuan).

Oreolalax omeimontis Hu et al., System. Index Amph.
China, p. 31.

Thirty five males, SVL 40-55 mm (mean 49.4);
weight 7-16 g (N = 25, mean 12.0). One female,
SVL 62 mm. One juvenile, SVL 33 mm. The fe-
male and seven males are from Mt. Emei, the rest
from Wa Shan.

All of the Emei specimens were caught by day
under rocks on the bank of a small (3 m) stream
at 1600 m. Most (21) of the Wa Shan frogs were
caught in broad-leaved forest at 1 360-1 530 m, six
were in mixed broad-leaved and coniferous forest
at 1290 m, and two were in a small clearing at
1400 m. All 28 adults were caught in or on the
banks of streams .5^4 m wide, the juvenile under
dead leaves 5 m from a small stream. Seven males
captured during the day were in water. Males call-
ing at night were in water (5), under large rocks
(10), on large rocks (4), or in gravel (2).

Megophrys omeimontis Liu

Oreolalax pingii (Liu)

Megophrys omeimontis Liu, 1950, Fieldiana: Zool.,
Mem., 2: 191. Type locality: Mt. Omei (= Emei),
Szechwan (= Sichuan); 3600 ft.

Fourteen males, SVL 47-57 mm (mean 51.1);
weight 8-16 g (mean 12.2). Four females, SVL 60-

Scutiger pingi Liu, 1943, J. West China Border Res.
Soc., ser. B, 14: 35. Type locality: Yenwot'ang (=
Yanwotang), Chaochiao (Zhaojue County), Szech-
wan (= Sichuan).

Oreolalax pingii Myers and Leviton, 1962, Copeia,
1962: 289.

FIELDIANA: ZOOLOGY

Seven males, SVL 41— 45 mm (mean 43.3). Five
females, SVL 37-46 (mean 41.8). Six juveniles,
SVL 26-35 mm. All from Qiliba.

These frogs were collected in chaparral at 2950-
3020 m. Only one was collected at night, in a
stream 4 m wide. The others were caught in or on
the banks of small (.5-2 m) streams, two at the
edge of a permanent pond, and one 8 m from a
stream. All except one were under cover, mainly
large rocks.

Oreolalax popei (Liu)

Scutiger popei Liu, 1947, Copeia, 1947: 125. Type
locality: Lungtung (= Longdong), Pao-hsing (=
Baoxing County), Sikang (now Sichuan).

Oreolalax popei Myers and Leviton, 1 962, Copeia,
1962: 289.

One female, SVL 63 mm. From Wa Shan in
broad-leaved forest at 1400 m. The frog was in a
small shallow depression about 5 m from a small
stream.

Oreolalax rugosus (Liu). Figure 4.

Scutiger rugosus Liu, 1 943, J. West China Border Res.

Soc., ser. B, 14: 37. Type locality: Yenwot'ang (=

Yanwotang), Chaochiao (= Zhaojue County),

Szechwan (= Sichuan).
Oreolalax rugosus Myers and Leviton, 1962, Copeia,

1962: 289.

One male, SVL 49 mm. Four females, SVL 47-
53 mm (mean 50.3). All from Qiliba at 2960-3020
m in chaparral. These specimens were collected in
or on the banks of small (.5-2 m) streams during
the day. Two were under large rocks and one under
a mat of moss.

Oreolalax schmidti (Liu)

Scutiger schmidti Liu, 1947, Copeia, 1947: 123. Type
locality: Mt. Omei (= Emei), Szechwan (= Sichuan),
7800 feet.

Oreolalax puxiongensis Liu and Fei, in Liu et al., 1979,
Acta Zootaxon. Sinica, 4: 84, figs. 7, 8. Type lo-
cality: Puxiong, Sichuan.

Twelve males, SVL 41-47 mm (mean 43.3);
weight 5-8 g (N = 9, mean 6.4). Two females, SVL
47-5 1 mm; weight 10 g (N = 2). Three males were
caught on Mt. Emei, and the rest of the specimens
at Wa Shan.

The Mt. Emei frogs were collected in coniferous
forest at 2280 m on the bank of a small (1.5 m)

permanent stream under a layer of moss. The 1 1
frogs from Wa Shan were caught in a coniferous
forest at 2520 m in or on the banks of small (1-2
m) permanent streams. Males were calling during
the day from under rocks or thick (20 cm) mats
of moss. With one exception, those not actually
in the stream (8 individuals) were within 20 cm
of the water's edge. Both females had ovulated;
one was under a large rock and the other under a
mat of moss at the stream margin.

The only differences between Oreolalax puxion-
gensis and O. schmidti, according to Liu and Fei
(in Liu et al., 1979), involved the number of spi-
nules on the backs of males and the color of the
ova. Males of puxiongensis were said to have more
clusters of black spinules on the back than males
of schmidti. However, Liu and Fei also noted vari-
ation in the spinosity of male paratypes of pux-
iongensis from Mianning Xian, about 75 km
southwest of the type locality. (Puxiong is about
75 km southwest of the type locality of schmidti.)
The nine males we collected at Wa Shan (40 km
west of Mt. Emei) show a wide range of variation
in spinosity, from few clusters of melanic spinules
on the back and almost none ventrally to clusters
covering most of the rear two-thirds of the back,
the dorsal surfaces of the hind limbs, the ventral
surface of the thigh, and the chin. One of us (GW)
found similar variation in 30 male schmidti from
Baoxing, 100 km northwest of Mt. Emei.

Ova of O. puxiongensis were described as hav-
ing purplish gray animal poles (Liu et al. 1979).
But this was the color of ova within the female.
One of us (GW) has collected egg masses of six
species of Oreolalax— major, omeimontis, pingi,
popei, rugosus, and schmidti— and all had white
or creamy white ova. Tian (1983) also reported
creamy white egg masses of O. chuanbeiensis.
Ovarian (and, therefore, immature) eggs examined
by us are grayish at the animal pole. We believe
that the eggs of the types of puxiongensis were
immature, ovarian ova. Besides the evidence from
other species in the genus, it would be surprising
indeed if eggs deposited under rocks, the pattern
for all Oreolalax, were pigmented.

Since in all other respects puxiongensis agrees
with schmidti, we believe them to be conspecific
and treat the former as a synonym.

Scutiger chintingensis Liu and Hu

Scutiger chintingensis Liu and Hu, 1 960, Scientia Sin-
ica, 9: 770. Type locality: Chinting (= Jinding), Mt.
Omei (= Emei), Szechwan (= Sichuan).

INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

Fig. 4. Oreolalax rugosus (Liu); snout-vent length 47 mm.

Three males, SVL 44—47 mm (mean 46.3);
weight 7 g (N = 3). One female, SVL 54 mm;
weight 1 1 g. All were collected on top of Wa Shan
at 2520 m in conifer-rhododendron forest. They
were in or on the banks of a small (2 m) stream.
The three out of water were under a root mat,
under a large rock, and in grass.

BUFONIDAE

Bufo andrewsi Schmidt

Bufo andrewsi Schmidt, 1925, Amer. Mus. Novit., no.
1 75, p. 1 . Type locality: Likiang (= Lijiang County),
Yunnan, 8500 feet.

Our samples from Wa Shan and Qiliba differ
enough to oblige us to present information on them
separately.

Wa Shan— Eleven males, SVL 63-78 mm (mean
70.6); weight 30-53 g (N = 10, mean 39.5). Eight
females, SVL 90-101 mm (mean 95.6); weight 84-
116 g (mean 97.8). Nine juveniles, SVL 20-46
mm; weight 1-10 g.

Half (14) of the toads were found in groves of
conifers, the rest roughly evenly distributed in
broad-leaved, mixed, and bamboo forests and in
clearings. Two were collected at 1550 m and the
remainder at 1 150-1300 m. Juveniles and adults
had similar distributions in these vegetation types
and altitudinal zones. Four were caught on stream
banks, two in a grassy marsh, and the rest more
than 5 m from water. All were at ground level on
soil and dead leaves except for seven under cover.

Qiliba— Nine males, SVL 61-68 mm (mean
64.8). Two females, SVL 72, 80 mm. One juvenile,
SVL 24 mm.

These toads were collected at 2950-3020 m in
a mixture of chaparral and wet grassland, the latter

FIELDIANA: ZOOLOGY

dotted with small ponds. Four were in a temporary
pond, four on the banks of a permanent pond, and
the others scattered. All but one of the terrestrial
toads were in grass.

The Qiliba and Wa Shan toads differ in several
respects. Those from Qiliba are smaller and have
relatively shorter legs and narrower heads (table
1 ). They have few or no melanic spinules dorsally,
and few of the ventral spinose tubercles are tipped
with melanin. Most of them have rounded glands
on the dorsal surface of the head. In contrast, the
Wa Shan toads have melanin-tipped spinose tu-
bercles on the belly and throat, as noted by Schmidt
(1927) in the description of andrewsi; the top of
the head is usually smooth.

At Wa Shan we collected many tadpoles (and
saw thousands) in moderate-sized (5-8 m) clear
rocky streams; none of the rows of denticles were
divided in 98% (356/363) of these larvae. At Qili-
ba we collected tadpoles in small (1-2 m) slow
streams (3 lots) and small permanent ponds (2
lots); 85% (63/74) have the inner upper row of
denticles divided.

Our samples agree for the most part with Ma-
tsui's ( 1 986) definition ofBufo andrewsi and differ
from B. gararizans popei Matsui in their well-de-
veloped webbing and smaller size (males < 88 mm,
females < 103 mm). Larvae from Wa Shan also
fit Matsui's definition of andrewsi: they live in
streams and lack divided toothrows. Larvae from
Qiliba are more like those of gargarizans popei.
Although we believe both samples represent an-
drewsi, the differences between them indicate that
a clear definition of andrewsi is still not in hand.

Table 1. Comparison of body proportions of Qiliba
and Wa Shan samples of adult male Bufo andrewsi. Too
few females were available to warrant analysis. Data are
given as thousandths of SVL.

N

Range

Median

Tibia/SVL*

Wa Shan

11

403-463

449

Qiliba

9

405^40
Head width/SVL**

432

Wa Shan

11

346-390

366

Qiliba

9

328-367

337

* Mann-Whitney U = 21, P < .05
** Mann- Whitney U = 10, P < .05

MICROHYLIDAE

Calluella yunnanensis Boulenger. Figure 5.

Calluella yunnanensis Boulenger, 1919, Ann. Mag.
Nat. Hist., (9), 3: 549. Type locality: Yunnanfu (=
Kunming), Yunnan.

Nineteen males, SVL 29-37 mm (mean 33.75).
All were collected at Qiliba in permanent (9) and
temporary (10) ponds at 2950 m. Males called
from the water's surface at night.

RANIDAE

Rana boulengeri Giinther

HYLIDAE

Hyla annectans (Jerdon)

Polypedates annectans Jerdon, 1870, Proc. Asiat. Soc.
Bengal, 1 870: 84. Type locality: Khasi Hills, Assam,
India.

Hyla annectans Giinther, 1875, Proc. Zool. Soc. Lon-
don, 1875: 576.

Four males, SVL 32-39 mm (mean 36.3); weight
3, 7 g (N = 2). Three from Wa Shan at 1 200-1 500
m, one from Xichang City at 1500 m. One of the
Wa Shan frogs was on the ground in a grove of
conifers; the other three were at ground level in
large clearings. We heard males calling from co-
nifers frequently at Wa Shan.

Rana boulengeri Giinther, 1 889, Ann. Mag. Nat. Hist.,
(6), 4: 222. Type locality: Ichang (= Yichang), Hupei
(= Hubei).

Nineteen mature males with spines on first two
or three fingers and on chest, SVL 84-136 mm
(mean 107.3); weight 88-370 g (N = 18, mean
203.3). Five males with spines on first finger only,
SVL 65-78 mm (mean 71.66); weight 44-72 g
(mean 56.25). Thirty-two mature females, SVL
80-122 mm (mean 102.4); weight 76-264 g (N =
26, mean 1 53.4). Eight subadult females, SVL 69-
79 mm (mean 73.40); weight 44-70 g (mean 56.0).
Seven unsexed juveniles, SVL 27-66 mm; weight
3-46 g. Seven from Qing Cheng Shan, 64 from Wa
Shan.

All from Qing Cheng Shan were caught at 990
m in water or on rocks at water's edge of a tree-
and shrub-lined creek (10 m wide) flowing through
cultivated fields. Upstream the creek passed

INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

Table 2. Comparison of body proportions in males
of Rana chaochiaoensis and R. japonica from Qiliba and
Wa Shan, Sichuan, respectively. Data are given as thou-
sandths of SVL.

N

Range Median

Head width/SVL*

chaochiaoensis

16

308^04 352

japonica

11

302-335 326
Head depth/SVL

chaochiaoensis

5

249-295 263

japonica

5

194-237 215
Tibia length/SVL**

chaochiaoensis

16

530-622 577

japonica

11

609-671 642

* Difference between species significant at P = .02;
Mann-Whitney U test.

** Difference between species significant at P < .01;
Mann-Whitney t/test.

through disturbed broad-leaved evergreen forest.
Altitudinal distribution at Wa Shan was: 1 1 50 m
(5), 1200 m (25), 1250 m (4), 1290 m (20), 1360
m (10). Most of the Wa Shan frogs were caught in
(35) or at water's edge of (18) rocky streams flow-
ing through coniferous, broad-leaved, or mixed
forests. Stream widths varied from .5 to 8 m; 25
frogs were caught on streams < 1 .5 m wide and 26
on streams 8 m wide. The three largest females
(1 1 1-122 mm) and two of the three largest males
(126, 136 mm) were at streams <1.5 m. Of the
25 females from Wa Shan, five had ovulated, 17
held mature ovarian eggs, and three were spent or
just beginning development of ova.

Rana chaochiaoensis Liu

Rana chaochiaoensis Liu, 1 946, J. West China Border
Res. Soc., Ser. B, 16: 7. Type locality: Chaochiao
City (= Zhaojue), Sikang (non Sichuan).

Sixteen males, SVL 49-59 mm (mean 52.8); tib-
ia length, head width, and head depth given in
Table 2. Two females, SVL 60, 62 mm; tibia length/
SVL .54, .55; head width/SVL .34, .35. All from
Qiliba.

These frogs were caught in a mixture of chap-
arral and grassland at 2950-3 140 m. Nine were in
a shallow pool in the floodplain of a small stream
meandering through grass; there were a dozen egg
masses in the pool. Three other frogs were cap-
tured in small streams, two in permanent ponds
and four in grass at the edge of ponds.

Liu and Hu (1959) treated this taxon as a sub-
species of Rana japonica, a status it maintained
in subsequent publications (e.g., Liu & Hu, 1961;
Hu et al., 1977; Tian & Jiang, 1986). But Zhao et
al. (1987) reported numerous differences in the
karyotypes of japonica and chaochiaoensis in-
volving centromere positions in five chromo-
somes and the positions of AgNORs and C-bands.
Differences between these taxa in larval denticles
are illustrated by Liu and Hu (1961), though they
still considered them as subspecies. In addition,
we have observed differences in coloration and
body proportions. Rana chaochiaoensis has dark
spots on the sides and a dark inverted V on the
back; japonica lacks both features. The dorsolat-
eral fold is thick in chaochiaoensis and thin in
japonica. Head width and depth relative to SVL
are greater in chaochiaoensis and tibia length greater
in japonica (table 2). Considering the many dif-
ferences between these two taxa, we believe they
should be treated as distinct species.

Rana daunchina Chang. Figure 6.

Rana musica Chang and Hsu, 1932, Contr. Biol. Lab.
Sci. Soc. China, Zool., 8: 1 57. Type locality: Hount-
sinping (= Hongchunping), Mt. Omei (= Emei), Si-
chuan.

Rana daunchina Chang, 1933, China J., 18: 209 (sub-
stitute name).

Three males, SVL 44—46 mm (mean 44.9),
weight 12 g (N = 2). One female, SVL 54 mm,
weight 1 5 g. All from Wa Shan in large clearings
at 1200 m. Two were in a permament pond and
two at the edge of a smaller pond.

We follow Hu (in Frost, 1985) in treating this
taxon as a species distinct from Rana adenopleura.
Liu (1950) and Liu and Hu (1961) noted the dif-
ferences between Fujian adenopleura and Sichuan
daunchina in breeding behavior, though still re-
garding the two as conspecific. Fujian and Sichuan
populations have distinct calls. Eggs of adeno-
pleura are laid as a globular cluster in ponds or
flooded rice fields, but those of daunchina are
placed in small burrows (see measurements in Liu,
1950) excavated by males next to ponds. We ob-
served one 4 cm burrow at the edge of a pond
measuring 1 00 x 70 x 20 cm and another 20 cm
from a pond measuring 300 x 300 x 40 cm.

Rana adenopleura has been reported from cen-
tral southern Sichuan (Shi & Zhao, 1982). One of
us (G W) has collected frogs of this group at those
Sichuan localities and in southeastern Guizhou

10

FIELDIANA: ZOOLOGY

Fig. 5. Calluella yunnanensis Boulenger, snout-vent length 33 mm.

and southeastern Hunan. In all those places the
frogs have the typical adenopleura call. In the only
area where egg masses were seen, Hunan, they
were laid directly in ponds.

As currently understood, daunchina is known
only from Mt. Emei and Wa Shan.

Rana japonica Gunther

Rana temporaria var. japonica Gunther, 1859, Cat.

Batr. Sal. Brit. Mus., p. 1 7. Type locality: Ningpo

(= Ningbo) and Chusan (= Zhou Shan Is.), Zhejiang,

and Japan.
Rana japonica Boulenger, 1879, Bull. Soc. Zool.

France, 1879: 190.

Two males, SVL 59, 62 mm; weight 20, 21 g.
Both from Wa Shan at 1 200 m, where they were
caught at the edge of a road passing through mixed
conifer and broad-leaved evergreen forest. In ad-
dition nine males (SVL 42-58 mm) and seven

females (SVL 43-62 mm) were caught by children
in and around rice fields in nearby Bin Ling at
980 m.

The relationship of this taxon to R. chao-
chiaoensis is discussed under that species.

Rana margaretae Liu

Rana margaretae Liu, 1950, Fieldiana: Zool., Mem.,
2: 303. Type locality: Panlungshan (= Mt. Panlong),
Kwanhsien (= Guan County), Szechwan (= Si-
chuan).

Thirty-one males, SVL 66-95 mm (mean 75.5);
weight 27-94 g (mean 40.7). Forty-nine females,
SVL 71-110 mm (mean 90.8); weight 32-102 g
(N = 44, mean 76.2). Four juveniles, SVL 39-60
mm; weight 6-19 g. Four are from Qing Chen
Shan, 80 from Wa Shan.

The four females from Qing Chen Shan were on
rocks at water's edge on a tree-lined, 10 m creek

INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

11

flowing through cultivated fields at 990 m. The
Wa Shan frogs were at 1 150-1360 m. A juvenile
was caught 30 m from a stream, but all the rest
were in water or on large rocks forming the banks
of streams flowing through broad-leaved, conif-
erous, and mixed forests. The majority (62) were
along a creek 8 m wide, and the rest along streams
1-5 m wide.

Rana shuchinae Liu

Rana shuchinae Liu, 1950, Fieldiana: Zool., Mem.,
2: 313. Type locality: Lolokou (= Jiefang Gou),
Chaochiaohsien (= Zhaojue County), Sikang (now
in Sichuan).

Eight males, SVL 3 1-34 mm (mean 32. 1). Three
females 27-39 mm (mean 34.3). All from Qiliba
at 2960 m in a mixture of chaparral and grassland.
We caught eight in or at the edge of permanent
ponds, two in amplexus in a temporary pond, and
one in an intermittent stream (.5 m).

Rana yunnanensis Anderson

Rana yunnanensis Anderson, 1879, Anta. Zool. Res.
Yunnan, p. 839. Type locality: Hotha (= Husa),
Yunnan.

Rana phrynoides Boulenger, 1917, Ann. Mag. Nat.
Hist., (8) 20: 413. Type locality: Tongchuan fu (=
Malong), Yunnan.

Rana (Paa) sichuanensis Dubois, 1986, Alytes, 5: 47.
Type locality: Ning Yuan Fu (= Xichang City), Si-
chuan.

Two males, SVL 75, 83 mm, 5 females, SVL
92-1 17 (mean 104.3), 5 juveniles SVL 46-79 mm.
Head width/SVL .38, .42 in males, .38-40 in fe-
males. All from Qiliba.

All were collected in a mixture of chaparral and
grassland at 2950 m. One juvenile was caught on
the bank of a small permanent pond, one female
in a temporary pond, and the rest in (6) or on the
banks (4) of small streams .5-1.0 m wide. Those
on the banks were in grass less than 30 cm from
the water's edge.

Liu and Hu (1961) believed Rana phrynoides
Boulenger and Rana yunnanensis Anderson were
conspecific. This opinion has been shared by sub-
sequent authors (e.g., Frost, 1985; Dubois, 1986).
Dubois (1986) based sichuanensis on two speci-
mens collected in 1914 in the mountains near
Xichang. Besides minor differences in body pro-
portions, R. sichuanensis was said by Dubois to

differ from yunnanensis in having a very distinct
tympanum, the webbing more excised between the
toes, the tips of the digits slightly but clearly di-
lated, the dorsum set with many short narrow lon-
gitudinal folds, and the skin fold along the outer
margin of the fifth toe ending proximally at the
level of the basal subarticular tubercle.

As Qiliba is in mountains near Xichang City,
our specimens are very nearly (if not actually) to-
potypes of sichuanensis. One-third (4) have short
narrow longitudinal folds on the back and sides;
all have many round wartlike tubercles, each
capped with a small whitish cone (presumably
equivalent to Dubois's "petites granulosites cor-
nees"). The skin fold along the fifth toe ends proxi-
mally, as Dubois describes, at the basal subarticu-
lar tubercle. The tips of the digits are slightly dilated.
However, the tympanum is obscure and not, as
Dubois says, very distinct.

In addition we have examined the following
specimens in the collection of the Chengdu Insti-
tute of Biology: (1)17 males from Longling Coun-
ty, Yunnan, 75 km from the type locality of yun-
nanensis; (2) 7 males from Jingdong and Xinping
counties, Yunnan, 300-400 km E of the type lo-
cality of yunnanensis and 220-300 km SW of the
type locality of phrynoides; (3) 1 2 males from Bin-
chuan, Dengchuan, and Lijiang, NW Yunnan, ap-
proximately equidistant among all three type lo-
calities; (4) 1 1 males from Huili and Huidong, S
Sichuan, approximately midway between the type
localities of phrynoides and sichuanensis; (5) 7
males from Zhaojue (= Chaochiao) 55 km E of
Xichang City near the type locality of sichuanen-
sis; (6) 10 males from Jiulong, Muli, and Luguhu
in the hills 1 10-150 km W of Xichang City; and
(7) 5 males from Weining, Guizhou, 175 km NE
of the type locality of phrynoides. These localities
are shown in Figure 7.

These frogs and the ones from Qiliba are clearly
conspecific. The only consistent geographic vari-
ation we can detect is in the dorsal skin. Frogs
from areas ( 1 ) and (2) have round, warty tubercles,
whereas those from areas (3)-(7) are like those
from Qiliba and have short skin folds or scattered
round spinose tubercles. Large males from south-
western (area 1) and southern (2) Yunnan have
black spines in two patches on the breast and on
the chin (as illustrated by Liu and Hu, 1961, fig.
48, p. 163); smaller males have them only on the
breast. Males from the other areas usually have
spines on the breast only; a few have weak spines
on the chin. Otherwise, there is individual varia-
tion in distinctness of tympanum and in the length

12

FIELDIANA: ZOOLOGY

Fig. 6. Rana daunchina Chang; snout-vent length 46 mm.

of the skin fold on the fifth toes. These observa-
tions confirm the conclusion of Liu and Hu ( 1 96 1 ),
though with yunnanensis as the prior name, and
lead us to place sichuanensis Dubois in the syn-
onymy of yunnanensis.

Amolops chunganensis (Pope)

Rana chunganensis Pope, 1929, Amer. Mus. Novit.,
no. 352, p. 3. Type locality: Kuatun, Chungan Hsien
(= Chongan County), Fujian.

Amolops chunganensis Matsui in Frost, 1985, Amph.
Species of the World, p. 453.

Nine males, SVL 34-38 mm (mean 36.3); weight
5-6 g (N - 8, mean 5. 1 ). One female, SVL 52 mm;
weight 1 6 g. All from Wa Shan.

All frogs were along streams flowing through
coniferous and mixed broad-leaved and conifer-
ous forests at 1 170 (6), 1200 (3), and 1400 m (1).
Only one was actually in water and the rest 1-5
m from water's edge on rocks (5), shrubs (2), a
small log (1), and soil (1). Males were actively

calling on several nights. The abdominal cavity of
the single female was filled with full-sized ova which
were not in gelatinous envelopes. The breeding
season may have been starting.

The geographic range of Amolops chunganensis
extends from Gansu Province in the northwest to
Fujian Province in the southeast; however, its dis-
tribution is apparently disjunct. One center lies in
northeastern Sichuan, southern Shaanxi, Gansu,
and north-central Sichuan; a second lies in western
Hunan and northern Guanxi; and a third in north-
western Fujian. Our specimens agree with the type
series, which we examined in the American Mu-
seum of Natural History.

Amolops loloensis (Liu)

Staurois loloensis Liu, 1950, Fieldiana: Zool., Mem.,
2: 353. Type locality: Lolokou (= Jiefanggao), Chao-
chiaohsien (= Zhaojue County), Sikang (now Si-
chuan).

Amolops loloensis Matsui, in Frost, 1985, Amph.
Species of the World, p. 455.

INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

13

25°

SICHUAN

JIULONG

ZHAOJUE

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105

Fig. 7. Southern Sichuan and adjacent provinces showing localities of samples of Rana yunnanensis. Type
localities: Hotha = R. yunnanensis Anderson; Malong = R. phrynoides Boulenger; Xichang (approximate) = R.
sichuanensis Dubois.

Two females, SVL 43, 61 mm. From Qiliba at
2950 m, 5 m from a stream bed. One was under
an overhanging rock and one on a large rock.

Amolops mantzorum (David)

Polypedates mantzorum David, 1871, Nouv. Arch.
Mus. Natl. Hist. Nat. Paris, 7, Bull.: 45. Type lo-
cality: Muping (= Baoxing), Szechwan (= Sichuan).

Amolops mantzorum Matsui, in Frost, 1985, Amph.
Species of the World, p. 455.

Five males, SVL 5 1-53 mm (mean 5 1 .6), weight

1 1-13 g (mean 12.0). Eleven females, SVL 60-70
mm (mean 65.7), weight 23-30 g (N = 8, mean
26.3). All from Wa Shan.

All were caught along stream courses, two in
water and the rest within 2 m of water's edge. Only
one was perched on a shrub, 12 on large rocks,
and one on gravel. Half were on an 8 m stream
and half on small streams .5-2 m wide. Five of
those at small streams were in forest clearings at
1400-1470 m. Ten were in broad-leaved ever-
green forest at 1250 (9) and 1400 m (1). One was
in mixed conifer and broad-leaved forest at 1250
m. We did not hear males calling. None of the
females was gravid; all were in middle stages of

14

FIELDIANA: ZOOLOGY

Fig. 8. Polypedates chenfui (Liu); snout-vent length 51 mm.

ovarian development with ova clearly not full-
sized.

RHACOPHORIDAE

Polypedates chenfui (Liu). Figure 8.

Rhacophorus chenfui Liu, 1 945, J. West China Border
Res. Soc., ser. B, 15: 35. Type locality: Hui-ting-
sze (= Hui-deng-si), Mt. Omei (= Emei), Szechwan
(Sichuan).

Polypedates chenfui Jiang, 1987, Acta Herpet. Sinica,
6: 37.

Thirteen males, SVL 31-39 mm (mean 35.2);
weight 3-4 g (N = 10, mean 3.6). Two females,
SVL 47, 5 1 mm; weight 9, 1 3 g. One juvenile, SVL
21 mm; weight 1 g. All from Wa Shan at 1200 m.

All but one of these frogs were caught in clear-
ings surrounded by broad-leaved and coniferous
forests. Ten were in herbs and low shrubs at the
margins of ponds, and of these, seven were caught
in a mixed chorus with P. omeimontis. Six were
at a distance from any body of water and four of
these were in vegetation 0.6-2.0 m above ground.

Polypedates dugritei David

Polypedates dugritei David, 1871, Nouv. Arch. Mus.
Natl. Hist. Nat. Paris, 7: 95. Type locality: Muping
(= Baoxing), Sichuan.

Eleven males, SVL 36—45 mm (mean 40.1);
weight 5-7 g (N = 8, mean 5.5). One female, SVL
43 mm; weight 6 g. One juvenile, SVL 21 mm.
Four from Qiliba, nine from Wa Shan.

The Qiliba frogs were at the margins of ponds
in mixed chaparral and grassland at 2960 m. One
was in grass, two on low shrubs, and the juvenile
under a rock. The Wa Shan frogs were caught at
2520 m, one in a small pond in a coniferous forest
and the others in grass around ponds in a large
marshy area surrounded by coniferous forest.

Polypedates omeimontis Stejneger

Polypedates omeimontis Stejneger, 1924, Occ. Pap.
Boston Soc. Nat. Hist., 5: 120. Type locality: Shin-
kai-sze (= Xin-kai-si), Mt. Omei (= Emei), Szech-
wan (= Sichuan).

INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

15

Thirteen males, SVL 54-67 mm (mean 59.5),
weight 1 1-24 g (N = 10, mean 14.1). Two from
Qing Chen Shan, 1 1 from Wa Shan.

The two from Qing Cheng Shan were in low
vegetation at the margin of a small (2 m) pond on
a farm at 990 m. One foam nest was attached to
grass at the pond's edge. The Wa Shan frogs were
collected at 1 200-1 260 m, five in shrubs and herbs
.05-1 m above ground at the edge of a 3 m pond
in a grassy clearing. Males called from this pond
on several nights, including one when male P.
chenfui also were calling. Five to 10 foam nests
were attached to shrubs overhanging this pond
throughout our stay at Wa Shan. We collected many
tadpoles of omeimontis from this pond. Five males
were caught in shrubs and small trees 1-2 m above
ground on the banks of streams in coniferous and
broad-leaved forest. Only one was found more
than 10 m from any body of water.

SAURIA
SCINCIDAE

Scincella monticola (Schmidt)

Leiolopisma monticola Schmidt, 1927, Bull. Amer.
Mus. Nat. Hist., 54: 496. Type locality: Snow
Mountain Village, Likiang (= Lijiang), Yunnan.

Scincella monticola Greer, 1974, Australian J. Zool.
Suppl. Ser., no. 31, p. 7.

Six males, SVL 37^49 mm (mean 44.0); tail
length/SVL 1.56, 1.68 (N = 2); weight 2-3 g (N =
5, mean 2.2); scale rows at midbody 22-24 (mean
23.5), scales parietal to thigh 53-64 (mean 59.0).
Four females, SVL 44-5 1 mm (mean 48.5); weight
1-3 g (mean 2.0), scale rows at midbody 22-24
(mean 23.5), scales parietal to thigh 58-62 (mean
59.8). One juvenile, SVL 36 mm; weight 1 g, scale
rows at midbody 24, scales parietal to thigh 58.
Supralabials 7 (6/7 in one, 7/8 in one); nuchals 2-
3; prefrontals separated in 10, meet at a point in
one (as in type); lamellae 10-13 (mean 1 1.5); scale
rows between lateral stripes 1/2 + 4 + 1/2. All
from Wa Shan.

Lizards were collected in groves of conifers (8),
mixed coniferous and broad-leaved forest (1), and
clearings (2) at 1200 (4), 1230-1270 (6), and 1400
m (1). Two were just below the surface of the soil,
five under rocks and dead branches, two on the
soil, one on a rock, and one on a dead branch. The
females had 1—4 enlarged eggs in the oviducts: 44

mm (1 and 2), 49 mm (2 and 2), 50 mm (3 and
4), 5 1 mm (2 and 3).

Scincella tsinlingensis (Hu and Zhao)

Leiolopisma tsinlingensis Hu and Zhao, 1966, Acta
Zool. Sinica, 18: 82. Type locality: Lao-hsien-cheng
(= Lao-xian-cheng), Chouchih Hsien (Zhouzhi
County), Shensi (= Shaanzi).

Scincella tsinlingensis Greer, 1974, Australian J. Zool.,
Suppl. Ser., no. 31, p. 7.

Three males, SVL 47—49 mm (mean 47.7), none
with complete tails, scale rows at midbody 28-30
(mean 28.7), scales parietal to thigh 78-85 (mean
82.3). Six females, SVL 46-59 mm (mean 53.5);
tail length/SVL 1.21 (N = 1); scale rows at mid-
body 26-28 (mean 27.0), scales parietal to thigh
78-80 (N = 4, mean 79.5). Two juveniles, SVL
30, 38 mm; scale rows 26. Supralabials 7 (6/7 in
one, 7/8 in one); nuchals 1-3; prefrontals in broad
contact (2), meet at a point (5), or separated (4);
lamellae 11-16 (mean 13.5); scale rows between
lateral stripes 1/2 + 4 + 1/2. All from Qiliba.

These skinks were caught at 2950 (7), 3020 (2),
and 3140 m (2) under small rocks and debris in
fields around farm buildings (7) or along a road
(4) through chaparral and grassland. Four females
had one or two developing embryos in each ovi-
duct.

The type locality is an area of broad-leaved for-
est at 1 800 m, unlike the environment at Qiliba.
However, two of us (EZ) and (GW) have also caught
this species at Hongyuan in northern Sichuan, a
region of grassland at about 3000 m.

Sphenomorphus indica (Gray)

Hinulia indica Gray, 1853, Ann. Mag. Nat. Hist., (2),

12: 389. Type locality: Himalayas.
Sphenomorphus indicus Schmidt, 1927, Bull. Amer.

Mus. Nat. Hist., 54: 494.

Eight males, SVL 64-77 mm (mean 72.5); tail
length/SVL 1.64-1.85 (N = 3, mean 1.78); weight
7-12 g (N = 4, mean 9.8); scale rows at midbody
34-36 (mean 34.3). Five females, SVL 69-96 mm
(mean 83.0); tail length/SVL 1.80 (N = 1); weight
1 3, 22 (N = 2), scale rows at midbody 34-36 (mean
35.7). Five juveniles, SVL 41-49 mm, weight 3 g
(N = 2). Supralabials 7; nuchals 0-1; prefrontals
in broad contact (1), meet at a point (1), separated
( 1 2); lamellae under fourth toe 1 4- 1 8 (mean 1 6; 1 );

16

FIELDIANA: ZOOLOGY

Fig. 9. Achalinus meiguensis Hu and Zhao; snout-vent length 362 mm.

scale rows between lateral stripes 1/2 + 6 + 1/2.
Six from Qing Cheng Shan, 12 from Wa Shan.

These lizards had limited altitudinal distribu-
tion at both sites: 990 m at Qing Cheng Shan and
1200-1250 m at Wa Shan. All except one were
collected in clearings, 6 under rocks, 3 on rocks,
and the rest on the ground. Four females contained
mature ova, three or four in each oviduct.

varies from left to right sides; maximum diameter
on left is .5 mm, on right 1.3 mm.

Both lizards were caught by residents in the vil-
lage of Bin Ling, 9 km east of the Wa Shan camp.
No details of capture site were available.

SERPENTES

ANGUIDAE

Ophisaurus harti Boulenger

Ophisaurus harti Boulenger, 1899, Proc. Zool. Soc.
London, 1899: 160. Type locality: Kuatun, Fukien
(= Fujian).

One male, SVL 177 mm, tail incomplete 167
mm, 18 scale rows across the back. One female,
SVL 221, tail incomplete 245 mm; weight 46 g;
1 7 scale rows across the back. Two scales between
rostral and prefrontal. Size of female's ear opening

AMBLYCEPHALIDAE
Pareas chinensis (Barbour)

Amblycephalus chinensis Barbour, 1912, Bull. Mus.
Comp. Zool., 40: 132. Type locality: Luluping (=
Lulubing, nr. Mt. Emei), Szechwan (= Sichuan).

Pareas chinensis Zhao, 1972, Zhongguo Shenei Jian-
suobian, p. 26.

One male, SVL 412 mm; tail length/total .34;
weight 18 g. ventrals 172; subcaudals 80. Five
females, SVL 410-51 1 mm; tail length/total .20-
.24 (mean .216); ventrals 172-182 (mean 177.0);

INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

17

subcaudals 65-72 (mean 69.2). Scale rows 15; su-
pralabials 6 (1), 6/8 (1), 7/8 (2), 8 (2); oculars 1-
1; temporals 2 + 3 (5), 2 + 3/4 (1). All from Wa
Shan.

Five were caught in clearings and one in mixed
coniferous and broad-leaved forest, 1 200-1 230 m.
Three were under mats of moss and one 20 cm
above ground in a small shrub. Vertical positions
of two were not recorded. Three females had one
egg in the left oviduct and two in the right.

COLUBRIDAE

Achalinus meiguensis Hu and Zhao. Figure 9.

Achalinus meiguensis Hu and Zhao, 1966, Acta Zoo-
taxon. Sinica, 3: 162. Type locality: Meigu, Sichuan.

Two males, SVL 362, 395 mm; tail length/total
.25, .21; weight 22, 28; ventrals 152, 155; subcau-
dals 56, 60. One female, SVL 405 mm; tail length/
total .16, weight 22 g; ventrals 165, subcaudals 44.
Two juvenile males, SVL 133, 138 mm; tail length/
total .21, .20; weight 2 g; ventrals 156, 159; sub-
caudals 6 1 , 62. One juvenile female, SVL 1 3 1 mm;
tail length/total .15; weight 2 g; ventrals 163; sub-
caudals 42. Scale rows 21-21-19 (5), 23-21-19 (1);
internasals absent; supralabials 3-2-1; oculars 0-
1, the postocular very small; temporals 2 + 2; anal
single. All from Wa Shan.

Snakes were caught at 1200 (2), 1300 (2), and
1400 m (2). Three were in coniferous forest, two
in broad-leaved evergreen forest, and one in a small
clearing. One was 1 5 cm below the surface at the
base of a rotting stump, one 5 cm below soil, one
under dead leaves, one under a rock, and one on
a stump. The female held two eggs in each oviduct.

These snakes, which represent only the second
record for this species, closely fit the original de-
scription, which was based on a specimen collected
in the 1930s. Two characters are diagnostic: a very
small postocular is present, internasals are absent.

Achalinus spinalis Peters

Achalinus spinalis Peters, 1869, Monatsber, Akad.
Wiss. Berlin, 1869: 146. Type locality: uncertain.

One juvenile, SVL 1 97 mm; tail length/total . 1 8;
weight 3 g; ventrals 1 54; subcaudals 60; scale rows
24-24-23; supralabials 3-2-1; oculars 0-0; tem-

porals 2 + 2. From Wa Shan at 1 230 m under thick
moss in a forest clearing

Amphiesma metusia sp. nov. Figure 10.

Holotype- Field No. 36982 (CIB), adult female;
Wa Shan, Hongya County, Sichuan Province,
China, alt. 1200 m; in forest clearing. 29 May
1987.

Paratypes— Four adult females, field no. 36849,
36874-5, 37514; two juveniles, 37271, 37302.
Same locality as holotype.

Diagnosis— This new form shares the following
characters with a number of Chinese species of
Amphiesma: a black stripe at side of belly formed
by a spot in lateral fourth of each ventral, a light
streak on temporal area separated from eye, 19
scale rows reducing to 1 7 near end of body and 8
supralabials. The new species differs from the oth-
er Chinese Amphiesma by having (1) a bold pat-
tern of alternating rows of black squares (not found
in any other Chinese Amphiesma); (2) 159-164
ventrals in females (differing from craspedogaster,
johannis, miyajimae, optata, parallela, and viba-
kari); (3) 72-85 subcaudals (differing from johan-
nis, miyajimae, optata, parallela, and vibakari);
and (4) dorsal head plates with a mottled pattern
lacking a pair of small, dark-edged, light spots on
the parietals (spots present in most species of Am-
phiesma).

Description (data on holotype in parentheses)—
Head elliptic, its anterior portion somewhat nar-
rower and posterior portion slightly wider than the
neck. Snout rounded. Rostral broader than high,
only its upper margin visible from above. Inter-
nasals squarish, a little narrower in front. Pre-
frontals larger than internasals. Frontal shield-
shaped, longer than the distance from it to tip of
snout, but shorter than suture between parietals.
Nostril rounded, in the center of nasal. Nasal long-
er than high, a groove from nostril to its upper
and lower margins, but not completely divided
into two shields. Loreal squarish, the loreal on
both sides of 36849 extending posteriorly into or-
bit below preocular. Eye moderate, pupil rounded.
One preocular and three postoculars. Anterior
temporals 2 (2) on both sides in four specimens,
1/2 in two specimens, 1/1 in one; posterior tem-
porals 1 in four specimens and 2 (2) in three. Su-
pralabials 8 or 7 (7/8), 3-2-3 in three specimens,
2-3-3 in two, 2-2-3 in one, and 2/3-2-3 in the
holotype. Infralabials 9 in the holotype, 9/10 in
one specimen, and 10 in five; first pair in contact

18

FIELDIANA: ZOOLOGY

Fig. 10. Amphiesma metusia sp. nov.; snout-vent length 530 mm.

behind mental; first 5 in contact with anterior chin
shields. Mental triangular. Chin shields in two
pairs, the posterior pair a little longer than the
anterior; posterior pair separated only at rear by
three small shields arranged in a triangle.

Dorsal scales in 19 rows on neck (18 in one
paratype), 19 at midbody, and 17 before vent;
prominently keeled except row 1 or rows 1-2. Ven-
trals 159-164 (163), mean 161.7; anal divided;
subcaudals 72-85 (83), mean 79.2 (N = 6, one
with broken tail). The second to fifth subcaudals
in a single row in 37271.

Maxillary teeth 23 or 24 (24) on each side; in-
creasing gradually to the rear; no diastema.

Total lengths of adults 705-885 mm (885); SVL
530-663 mm (663); tail length .24-.27 of total
length (.25); weights 49-1 34 g ( 1 34). Total lengths
of juveniles 246 (incomplete tail), 387 mm; SVL
221, 281 mm; weights 3, 9 g.

Coloration a dark checkered pattern and a poor-
ly defined, usually interrupted medium brown
stripe on scale rows 6-7; ventrally red with narrow

black stripes formed by a mark near both lateral
corners of each ventral; an oblique black stripe
from rear of eye across last supralabials and con-
tinuing on to trunk; neck with a light band 3—4
scales wide separated from eye and forming an-
terior end of lateral stripe; top of head dark olive,
lighter on snout with black mottling but no paired,
light parietal spots; upper lip bright light yellowish
green with black sutures between the supralabials;
no markings on head ventrally.

Etymology— Met usia from metousia (Gr., part-
nership), in recognition of the joint Chengdu In-
stitute-Field Museum project.

Comparisons— The combination of conspicu-
ous ventral stripes and 19 scale rows in Amphies-
ma metusia resembles 10 species of the genus
known from China, including four reported from
Sichuan: craspedogaster, octolineata, optata, and
johannis. The first three have a pair of small light
parietal spots (as do the other six species alluded
to), setting them off from the new species. None
of the four has a checkered pattern, such as typifies

INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

19

metusia. Besides the coloration differences, metu-
sia is distinguished from craspedogaster in ventral
counts (138-157 in craspedogaster) and in having
the nasal scale undivided.

Apparently, metusia is also a larger species. Pope
(1935) said that the three largest females of 60
craspedogaster measured 478-490 mm SVL, which
is smaller than any of our adult females. We have
examined the Sichuan craspedogaster Pope listed.
They all resemble craspedogaster in coloration and
counts (ventrals < 150). Amphiesma octolineata
has, as its name indicates, a distinctly striped pat-
tern. Amphiesma optata has bold transverse light
bands and much higher subcaudal counts (95-1 12).
The only species agreeing with metusia in lacking
parietal spots isjohannis, which differs from metu-
sia in lacking a checkered pattern, in ventral ( 1 65-
1 78) and subcaudal (84-89) counts, and in having
the internasals strongly narrowed in front.

Five of these snakes were collected at 1200 m,
one at 1250 m, and one at 1470 m. Three were in
clearings and four in coniferous forest or mixed
broad-leaved and coniferous forest. One juvenile
was next to a permanent pond swarming with
Polypedates tadpoles, two adults were on the banks
of streams, and the rest away from bodies of water.
Both juveniles were off the ground in low vege-
tation. Four females contained 5-7 large oviducal
eggs; the holotype held seven.

on bare soil or dead leaves. Two females had one
enlarged egg in each ovary.

Plagiopholis styani (Boulenger)

Trirhinopholis styani Boulenger, 1 899, Proc. Zool. Soc.

London, 1899: 164. Type locality: Kuatun, Fukien

(= Fujian).
Plagiopholis styani Zhao, 1972, Zhongguo Shenei

Jiansuobian, p. 23.

One adult male SVL 273 mm, one juvenile male
SVL 109 mm tail length/total .15 (N = 2); weight
of juvenile 2 g; ventrals 109, 1 10; subcaudals 27,
29. Two adult females SVL 298, 313 mm, 2 ju-
venile females SVL 101, 110 mm; tail length/total
. 1 1— .13 (mean .121); weight of adults 22, 30 g, of
juvenile 2 g; ventrals 109-118 (mean 115); sub-
caudals 23-26 (mean 24.8). Scale rows 15; supra-
labials 2-2-2; oculars 1-2 (5), 1-1/2(1); temporals
2 + 2 (5), 1+2 (1). All from Wa Shan.

Four were collected in small clearings and one
in coniferous forest, all under moss of floor debris
at 1200-1230 m. One female had two eggs in each
oviduct, the other female two in the left and four
in the right.

Rhabdophis nuchalis pentasupralabialis

Jiang and Zhao

Amphiesma sauteri (Boulenger)

Tropidonotus sauteri Boulenger, 1 909, Ann. Mag. Nat.

Hist., (8), 4: 495. Type locality: Kosempo, Formosa.
Amphiesma sauteri Malnate, 1960, Proc. Acad. Nat.

Sci. Philadelphia, 112: 51.

One juvenile and three adult males (juvenile not
measured before preservation), SVL 178-295 mm;
tail length/total .37-39 (mean .38); weight of ju-
venile 1, of adults 4-13 g (mean of adults 7.7);
ventrals 132-138 (mean 134.8); subcaudals 74-80
(mean 78.3). One juvenile and three adult females
(one adult and the juvenile not measured or
weighed before preservation), SVL 340, 360 mm;
tail length/total .22 (N = 1); weight 16, 23 g; ven-
trals 134-139 (mean 135.2); subcaudals 72 (N -
2). Scale rows 17 throughout; supralabials 2-2-3;
oculars 1-2 or 1-3, 2-3 in one; temporals 1 +2 (6),
2 + 2 (2). All from Wa Shan.

All were collected at 1 200 m, five in coniferous
forest and three in large forest clearings. Four were
under cover (moss, dead leaves, or rock) and four

Tropidonotus nuchalis Boulenger, 1891, Ann. Mag.
Nat. Hist., (6) 7: 281. Type locality: Ichang (= Yi-
chang), Hupeh (= Hubei).

Rhabdophis nuchalis pentasupralabialis Jiang and
Zhao, 1983, Acta Herpet. Sinica, 2: 60. Type lo-
cality: Jiulong County, Sichuan.

Two adult and 3 juvenile males, SVL 327-486,
1 37-250 mm; tail length/total . 1 9-.24 (mean .211);
weight of adults 14-44 g, of juveniles 3-9 (mean
6.3) g; ventrals 148-154 (mean 150.6); subcaudals
55-59 (mean 58.0). Five adult and 3 juvenile fe-
males, SVL 342-523 mm, 138-241 mm; tail
length/total .17-18 (mean .172); weight of adults
1 7-55 g (mean 39.8), of juveniles 2-7 g (mean 5.0);
ventrals 148-1 57 (mean 1 5 1 .8); subcaudals 43-49
(mean 46.0). Scale rows 15; supralabials 2-2-1 (9),
2/3-2-1 (1), 2-1/2-1 (2), 2-2-1/2 (1); oculars 1-3
(9), 1-2/3 (2), 1/2-3 (1), 2-3 (1); temporals 1 + 1
(8), 1 +2 (5). All from Wa Shan.

These snakes were collected in a very narrow
altitudinal band, 1200-1250 m, considerably be-
low the elevation (2750 m) of the type locality of
this subspecies. Three were caught within forests,

20

FIELDIANA: ZOOLOGY

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INGER ET AL.: AMPHIBIANS AND REPTILES FROM SICHUAN

21

Table 4. Distribution of stream-breeding frogs along
streams of various widths at Wa Shan, Sichuan; figures
indicate number of individuals.

Stream width (m)

Species

0.5-2

3-5 8

Leptolalax pelodytoides

28

1 0

Megophrys omeimontis

11

1 2

Oreolalax omeimontis

23

5 0

Oreolalax schmidti

11

0 0

Scutiger chintingensis

4

0 0

Bufo andrewsi

2

0 2

Amolops chunganensis

1

0 9

Amolops mantzorum

8

0 8

Rana boulengeri

25

8 29

Rana margaretae

3

4 70

620 mm, 210 mm (all measured after preserva-
tion); tail length/total . 14-. 1 5 (mean .146); weight
of adults (before preservation) 220-242 g (mean
228.7); ventrals 141-143 (mean 142.25); subcau-
dals 41—45 (mean 42.5). Scale rows at neck 23-
29, at midbody 21-23, before vent 18-19; supra-
labials 9 (3) or 10 (4); 7-10 scales between supra-
oculars. All from Wa Shan.

Snakes were caught in clearings (5) and conif-
erous forest (2) at 1 200 m. Three were under floor
debris, two on surface of soil, and one each on and
under rocks. Two females were gravid, one with
seven and one with nine oviductal eggs.

one each in coniferous, broad-leaved, and mixed
forest. The rest were in clearings. Five were under
mats of moss, one under dead leaves, and one
under a rock. Six were on bare soil. Two females
had four and seven elongate, oviductal eggs.

VIPERIDAE

Trimeresurus jerdoni Giinther

Tri meresurus jerdoni Giinther, 1875, Proc. Zool. Soc.
London, 1875: 233. Type locality: Khasya(= Khasi
Hills, Assam).

One male, SVL 652 mm; tail length/total .17;
weight 102 g; ventrals 174; subcaudals 64. One
female, SVL (after preservation) 590 mm; tail
length/total .14; ventrals 177; subcaudals 57. Scale
rows 21-21-17, 23-21-17; supralabials 7; scales
between supraoculars 7. From Wa Shan.

Both were caught in clearings, one at 1200 m
and one at 1470 m. One was found under debris
and one draped over a low plant. The female had
three oviductal eggs on one side and four on the
other; none of the eggs had developing embryos.

Trimeresurus nl kola Giinther

Trimeresurus monticola Giinther, 1864, Rept. Brit.
India, p. 388. Type locality: Nepal.

Three males, SVL 425, 445 mm (one badly
damaged); tail length/total .18 (N = 2); weight 72
g (N = 1); ventrals 138, 142; subcaudals 49, 53.
Three adult and one juvenile females, SVL 610-

Summary of Habitat Distribution of
Anurans at Wa Shan

Our data on this topic can be used to give a
picture of ecological distribution of this assem-
blage only within a narrow interval of time. For
the majority of frogs that interval fell during or
close to the breeding season, so that our data refer
largely to breeding sites. Only one anuran for which
sample size was at least 10 was collected mainly
in nonaquatic areas— Bufo andrewsi (table 3).

Most of the Wa Shan frogs are stream breeders
(table 3), the five pelobatids, Bufo andrewsi, the
two Amolops, Rana boulengeri, and Rana mar-
garetae. In the hilly terrain of Wa Shan, streams
have rocky beds and banks. Therefore, the heavy
use of rocks as perch sites by these stream breeders
shown in the table is probably a matter of avail-
ability rather than selectivity. Nonetheless, perch
site varied among species. For example, Rana
margaretae and Amolops mantzorum used rocks
on the banks to a significantly greater extent than
other species (P < .01 in both cases; see Materials
and Methods). Only one species, Leptolalax pe-
lodytoides, commonly perched above ground on
vegetation. Oreolalax omeimontis, O. schmidti, and
Scutiger chintingensis, unlike the other species,
were found mainly under rocks; males commonly
called from that position. Although some species,
such as Rana boulengeri, R. margaretae, and the
two Amolops, appear to remain along stream
courses throughout the live cycle, only R. boulen-
geri among them was seen in water more often
than not (P < .01)

There were also differences among stream-
breeding species in terms of the widths of streams
exploited (table 4). The pelobatids as a group were
rarely seen at the widest stream, whereas the ranids

22

FIELDIANA: ZOOLOGY

Table 5. Distribution of size classes of Rana bou-
lengeri along streams of various widths at Wa Shan,
Sichuan.

Stream widths (m)

Snout -vent

0.5-2

3-5

8

length (mm)

No.

of individuals

<61

1

0

3

61-75

2

0

8

76-90

3

2

4

91-105

8

4

9

106-136

11

2

5

made heavy use of that habitat. The contrast be-
tween the distributions of Leptolalax pelodytoides
and Rana margaretae emphasize this point. With-
in the Ranidae, the difference between Rana
boulengeri and R. margaretae seen in Table 4 is
statistically significant (2x3 contingency table,
chi-square 30.53, P < .01). This difference is not
related to the fact that the sample of boulengeri
includes a much larger proportion of juveniles and
small adults. As Table 5 shows, all size groups of
Rana boulengeri occurred in both the narrowest
and widest streams searched.

The few pond breeders fall into two groups, one
comprising the three arboreal species ofPolypeda-
tes and the other two terrestrial ranids, Rana daun-
china and R. japonica. We observed calling males
and nest holes of R. daunchina at a pond where
P. omeimontis and P. chenfui were breeding. We
collected R. japonica near a roadside ditch that
contained tadpoles of that species. Polypedates du-
gritei is isolated from its two congeners because
of its restriction to high elevations; we found it
only on the top of Wa Shan, at 2520 m and the
others only below 1300 m.

We conclude from our observations that during
the breeding season there is a modest amount of
ecological segregation within related groups of
species. Among the pelobatids, Oreolalax schmidti
and Scutiger chintingensis were restricted to high
elevations (2520 m) and the others to below 1450
m. We found the two high altitude species in sep-
arate streams, but because our samples are small
and because the flattish top of Wa Shan is a net-
work of small streams, we are not certain about
their spatial isolation. Three of the pelobatids from
lower elevations— Oreolalax omeimontis, Lepto-
lalax pelodytoides, and Megophrys omeimontis—
were caught along the same small streams and
during the same collecting forays, although they
differ slightly in perching sites (table 3). We have

already noted differences between Rana boulen-
geri and R. margaretae in perch sites (table 3) and
relative abundances along small and large streams
(table 4). Nonetheless, both were caught along the
same segments of large streams during every col-
lecting foray at those places.

Acknowledgment

We wish to express our deep gratitude to the
people of the village of Bin Ling, who not only
helped us collect at Wa Shan but also were kind
to us in many ways. We are also indebted to Xiao
Fu, who provided shelter and much-needed hot
tea at Qiliba. Tan An-ming, Chengdu Institute of
Biology, participated in all the fieldwork, and we
are grateful to him for his efforts. Figure 1 was
prepared by Clara Richardson Simpson, Field Mu-
seum of Natural History. This project received
support from National Science Foundation grant
BSR 8602046 and National Geographic Society
grant 3418-86.

Literature Cited

Dubois, Alain. 1981. Notes sur la systematic et la
repartition des amphibiens anoures de Chine et des
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24

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